Siphonaria denticulata Quoy & Gaimard, 1833

Siphonaria denticulata Quoy & Gaimard, 1833 View in CoL

( Figs 19G–L, R View FIGURE 19 , 20D–H View FIGURE 20 )

Siphonaria denticulata Quoy & Gaimard 1833: 340 View in CoL , pl. 25; figs 19– 20 (type locality: Nouvelle-Hollande, port Western [Western Port, French Island, Vic, Australia]).— Lamarck 1836: 559, 1839: 206; Reeve 1842: pl. 138, fig. 2; Catlow & Reeve 1845: 100; Menke 1852: 38; H. Adams & A. Adams, 1855 (in 1853–58): 271; Reeve 1856 pl. 1, species 4; Hanley 1858b: 151; H. Adams & A. Adams 1863: 271; Angas, 1867: 232; Paetel 1873: 117; 1875: 92; Tenison Woods 1878b: 99; Paetel 1883: 178; Etheridge 1887: 29; Henn & Brazier 1894: 178; Brazier 1889; 25; Paetel 1889: 428; Whitelegge 1889: 117; Adcock 1893: 11; Hedley 1909; 369; 1918: M96; McAlpine 1952: 41; Ewers 1961; Galindo 1977: 416; Crease 1980: 38; Jenkins 1981: 2; Hochlowski et al. 1983: 7413; Quinn 1983: 81; Trew 1983: 2; Jenkins 1983: 29; 1984: 113; Hochlowski & Faulkner 1984: 3839; Short & Potter 1987: 122, pl. 60, fig. 3; Jansen 1995: 89; Davey 1998: 118, text-fig.; Grove et al. 2006: 60; White & Dayrat 2012: 62; Colgan & da Costa 2013: 74.

Siphonaria denticula Anton 1838: 26 (invalid; incorrect subsequent spelling of S. denticulata View in CoL ).

Siphonaria atra View in CoL — Angas 1867: 233; Whitelegge 1889: 117.

Siphonaria cochlearformis Whitelegge 1889: 117 ; Hedley 1918: M96 (invalid; incorrect subsequent spelling, not S. cochleariformis Reeve, 1856 View in CoL ).

Ellsiphon marza Iredale 1940: 438 View in CoL pl. 34, figs 1, 2 (type locality: Keppel Bay, Queensland).— McAlpine 1952: 42; White & Dayrat 2012: 57, 60, 63.

Siphonaria scabra View in CoL — Iredale 1940: 438 (not S. scabra Reeve, 1856 View in CoL ).

Siphonaria marza View in CoL — Hubendick 1946: 61, pl. 5, fig. 27, 28; Galindo 1977: 416.

Siphonaria (Ductosiphonaria) diemenensis var. denticulata View in CoL — Hubendick 1946: 38, pl. 2 fig. 17.

Ellsiphon denticulatus — McAlpine 1952: 41, fig. 1; Iredale & McMichael 1962: 82.

Siphonaria (Siphonaria) currumbinensis Hubendick 1955: 126– 136 View in CoL , figs 4–8, pl. 1, figs 1–6 (type locality: Currumbin, Qld, Australia).— White & Dayrat 2012: 62, 70.

Ductosiphonaria denticulata — Cotton 1959: 411.

Siphonaria alternicosta View in CoL — Cotton 1959: 411 (not S. alternicosta Potiez & Michaud, 1838 View in CoL ).

Siphonaria View in CoL ‘ atra View in CoL group, unit 33’— Dayrat et al. 2014: 264, fig. 5 J (not S. atra Quoy & Gaimard, 1833 View in CoL ).

Material examined. Type material. Lectotype of Siphonaria denticulata Quoy & Gaimard, 1833 , present designation, from ‘partie sud de la Nouvelle-Hollande, au port Western, et probablement aussi au port du Roi-Georges’ [southern part of Australia, at Western Port, Vic, Australia, and probably also at King George Harbour ; see comments below] ( MNHN IM 2000-35951 , Fig. 19G View FIGURE 19 ). Two paralectotypes, same data as lectotype ( MNHN IM 2000-5062 ).

Probable holotype of Ellsiphon marza Iredale, 1940 from Keppel Bay [Qld, Australia]; coll. H. Bernhard, 1940 ( AM C.103715 , Fig. 19K View FIGURE 19 ). Probable paratypes of Ellsiphon marza Iredale, 1940 . Similar data to holotype label (refer Taxonomic remarks) ( MV F13843 , 3 d); Tampian Heads, Yeppoon , Keppel Bay ; coll. H. Bernhard Dec 1938 ( AM C.58529 , 26 d); Four probable paratypes; coll. 1940 ( AM C.106867 ).

Holotype of Siphonaria currumbinensis Hubendick, 1955 from Currumbin [28°07.83′S, 153°29.98’E, Qld, Australia]; coll. R. Endean & R. Kenny, 20-6-1952 ( MV F15562 , Fig. 19I View FIGURE 19 ). GoogleMaps Probable paratypes, same data as holotype ( MV F13953 , 2 d, 3 p; MV F13948 , 2 d, 3 p). GoogleMaps

Other, non-type material. Australia; Qld: Wreck Pt Yeppoon, 23°08.736’S, 150°45.865’E, Q08-4 ( AM C.585471 3p, C.584743 p [SK030 protoconch B7 jaw], C.585133 p [M081], C.585134 p [SK073]); GoogleMaps Double Head Yeppoon, 23°09.908’S, 150°47.638’E, Q08-3 ( AM C.585561 3p); GoogleMaps Bagara Hervey Bay , 24°49.180’S, 152°28.011’E, Q06-1 ( AM C.585961 p); GoogleMaps Alexandra Mooloolaba, 26°40.342’S, 153°06.822’E, Q04-1 ( AM C.585383 10p); GoogleMaps Nth Reef Scarborough, 27°11.432’S, 153°06.755’E, Q03-5 ( AM C.585560 4p); GoogleMaps Drury Pt Scarborough, 27°11.836’S, 153°06.955’E, Q03-7 ( AM C.585408 10+p); GoogleMaps Scarborough Pt Scarborough , 27°12.168’S, 153°06.980’E, Q03-8 ( AM C.585638 5p); GoogleMaps Currumbin Point, 28°07.523’S, 153°29.203’E, Q02-2 ( AM C.585536 20+p, C.585122 p [M073], C.585123 p [M074], C.585124 p [M183], C.585125 p [M184]) GoogleMaps . NSW: Brunswick Heads, 28°32.297’S, 153°33.444’E, NSW12- 1 ( AM C.585480 16p, C.585066 p [M077], C.585067 p [M079]); GoogleMaps Evans River at South Evans Head , 29°6.82’S, 153°25.93’E ( AM C.339263 p); GoogleMaps Wooli Wooli River at Wooli , 29°53.27’S, 153°15.98’E ( AM C.339285 4p; AM C.339287 2p); GoogleMaps Redbank (Corindi) River at Red Rock , 29°59.03’S, 153°13.55’E ( AM C.339288 3p); GoogleMaps Urunga mangroves on S side of town just beyond footbridge, 30°29.92’S, 153°1.16’E ( AM C.355024 8p); GoogleMaps Camden Haven Inlet , 31°38.37’S, 152°49.63’E ( AM C.339289 p); GoogleMaps Fingal Bay nr Port Stephens , 32°44.087’S, 152°10.402’E, NSW09-2( AMC. 5856837 p), GoogleMaps 32°44.990’S, 152°10.481’E, NSW09-1 ( AM C.585065 p [M151]); GoogleMaps Bolton Point Lake Macquarie , 33°0.389’S, 151°36.889’E, NSW08-3 ( AM C.585737 2p, C.585058 p [M452]; C.585059 p [M453]); GoogleMaps Bateau Bay Beach , 33°23.003’S, 151°29.197’E, NSW08- 6 ( AM C.595940 p [SK520); GoogleMaps Crackneck , Bateau Bay , 33°23.316’S, 151°29.139’E, NSW08-5 ( AM C.585063 p [SK295], C.585064 p [SK296]); GoogleMaps Broken Head Terrigal , 33°26.796’S, 151°27.030’E, NSW08-1 ( AM C.585528 20+p, C.585050 p [M001]); GoogleMaps Terrigal The Skillion , 33°27.008’S, 151°27.122’E, NSW08-2 ( AM C.585558 3p, C.585053 p [M226], C.585054 p [M225]); GoogleMaps Brisbane , WAter, 33°29.82’S, 151°18.18’E ( AM C.355025 8p); GoogleMaps Fairlight Sydney Harbour , 33°47.971’S, 151°15.990’E, NSW06-4 ( AM C.585590 4p); GoogleMaps Fairlight , North Harbour , 33°47.986’S, 151°16.837’E, NSW06-1 ( AM C.585589 6p); GoogleMaps 100m NW of Spit Bridge , Port Jackson , 33°48.210’S, 151°14.664’E ( AM C.546770 2p); GoogleMaps Spit Bridge , Port Jackson , 33°48.270’S, 151°14.520’E ( AM C.546768 p); GoogleMaps Wy-ar-gine Point Balmoral, 33°49.159’S, 151°15.195’E, NSW06-5 ( AM C.585476 17p, C.585043 p [M158], C.585044 p [M159], C.585045 p [M160], C.585046 p [M161], C.595923 p [SK031]); GoogleMaps Laings Point Sydney Harbour, 33°50.419’S, 151°16.638’E, NSW06-3 ( AM C.585589 5p, C.585034 p [M193], C.585041 p [M165], C.585042 p [M167]); GoogleMaps Bombo Kiama 34°39.232’S, 150°51.649’E, NSW03-1 ( AM C.585403 10+p, C.585333 p [SK043]); GoogleMaps Murunna Point Camel Head, 36°22.720’S, 150°04.766’E, NSW02-1 ( AM C.585401 10+p) GoogleMaps . Victoria: Bastion Head Mallacoota , 37°34.429’S, 149°45.927’E, V09-1 ( AM C.585787 5p, C.585294 p [M194], C.585295 p [M195]); GoogleMaps Cape Conran, 37°48.798’S, 148°43.608’E, V08-2 ( AM C.585292 p [M197]) GoogleMaps . Tas: T01-1 S of Granite Point Bridport , 40°59.739’S, 147°23.468’E ( AM C.584834p [M173]) GoogleMaps .

Taxonomic remarks. The largest syntype with the clearest external sculpture ( Fig. 19G View FIGURE 19 ) is herein designated as the lectotype of S. denticulata for the stabilisation of the name and to ensure the unambiguous identity of this taxon (Art. 74.1 of the Code). Our delineation of this species is based on comparative analyses of the morpho-anatomy and mitochondrial genetics of freshly collected topotypes of S. denticulata ( AM C.585303 [SK043], C.585044 [M159], C.585053 [M226]), Fig. 19H View FIGURE 19 ), Ellsiphon marza ( AM C.584743 [SK030], C.585133 [M081], Fig. 19L View FIGURE 19 ) and S. currumbinensis ( AM C.585123 [M074], Fig. 19J View FIGURE 19 ) and geographic series of additional specimens (Table S1). Quoy & Gaimard’s (1833: 340) statement ‘probably also at King George Harbour’ is not considered as part of the type locality. While there is also a King George Harbour in Tas, the King George Harbour referred to by Quoy & Gaimard (1833) is probably that near Albany, WA. However, we have not found this species in WA. The type locality for E. marza is Keppel Bay and Caloundra as stated in the original description, yet some paratypes are from additional sites (Tampian Heads and Yeppoon). Noosa, Qld is mentioned by Hubendick as a site for ‘ S. currumbinensis ’ The locality on the holotype label ( MV F15562) is misspelled as Corrumbin (sic Currumbin) and the name is misspelled as S. corrumbinensis . Hutton (1873: 55) stated that S. denticulata and S. diemenensis occur in NZ; however, this is likely based on misidentifications of S. australis and/or S. propria .

External morphology ( Fig. 19R View FIGURE 19 ). Foot sole smooth, evenly pale yellow grey; foot wall pale yellow grey with evenly spread white subepithelial pustules becoming more vivid and dense around pneumostomal lobe; foot wall shows black blotches; fringing mantle translucent, covers shell mantle, outer edge lobed, thickened, with black pigmented blotches aligning with rib interstices; pneumostomallobewithinmantlebetweentherightADMs, closes the pneumostomal and anal openings at the mantle edge; two small black epithelial eye spots centralised on two thick brownish yellow centrally touching cephalic folds; genital pore inconspicuous, located on foot wall to right anterior of right cephalic fold.

Shell ( Figs 19G–L View FIGURE 19 ; Table S9). Medium to large sized (max sl mean = 24.46 mm, SD = 2.04 mm, n = 20), ovate; height medium; apex offset sightly posterior and central (usually eroded), apical sides strongly convex, protoconch direction undetermined, shell whorl dextral; growth striae prominent in bands, shell thickness thick; rib count (mean = 42.6, SD = 4.5, n = 20), primary ribs pale white, fairly straight, increasingly raised and protrude beyond shell lip to unevenly scallop and corrugate the edge; 1–2 interspersed pale white finer secondary ribs, rib interstices darker; paired primary ribs on siphonal ridge, no more prominent than other primary ribs. Interior shell margin dark brown to tan, white rays align on shell margin under primary/secondary ribs, siphonal groove distinct, same colour as shell edge, points to right anterior; spatula dark chocolate brown to mottled tan even whitish; ADM scar distinct, CMS straight, paler than shell lip; thickening of shell lip translucent, infills and reduces lip scalloping, spatula becomes whitened. The shells of S. denticulata , S. scabra , S. pravitas sp. nov. and S. diemenensis may appear similar and may be misidentified; external ribbing is similar. Differences exist in the sharpness and intensity of primary ribs and rib interstice colouration. In addition, the spatula colouration differs; S. denticulata and S. scabra dark chocolate to tan-brown, S. diemenensis and S. pravitas sp. nov. are more golden brown.

Reproductive system ( Figs 20D, F, G; n View FIGURE 20 = 3). Positioned within coelom under the respiratory cavity, over foot muscle, against right side of foot wall; epiphallic parts positioned between BM and RAM; single GP inconspicuous, opening from GA through foot wall posterior to right cephalic fold. Join of AO, GA and ED distinct, all whitish fibrous muscular tissue; AO elongated, bluntly pointed, folded, larger than GA, thicker than ED; ED relatively long, thick and twisted; EG broad, lobed, bluntly pointed with two curved, broad, flagellum (F1 and F2; F2 twice length of F1); BD and CD connect close together into GA between ED, AO and GP; BD noticeably longer and thinner than CD with a prominent twisted loop immediately before BC (no distal loop or MA), both ducts smooth, featureless, pass together through RAM connecting into soft curved folds of MG, BD dorsal to CD; BC large, bulbous, thin test, internally embedded into folds of MG; HD short thick, coiled, links AG to elongated narrow yellowish granulated HD; HG larger than AG, MG and AG folded, soft white tissue; SV embedded on left side of AG under BC; AG and HG sides match curvature of inner foot wall at right posterior quarter of coelom.

Spermatophore ( Figs 20E, H View FIGURE 20 ). Head section over half of total length (head length = 8.08 ± 0.42 mm; ~ 43% of SPM length; head width = 119 ± 3 μm; flagellum width = 33 ± 1 μm, n = 3), cylindrically elongate, tip bluntly rounded, containing a white gelatinous mass; tapers quickly into consistently filamentous transparent flagellum; both sections smooth, translucent, test thin, featureless. multiple tightly coiled SPM in bursa, embedded in brown gelatinous mass. Seven and two SPM found in BC of two specimens [SK043, SK030].

Radula and jaw ( Figs 83E–H View FIGURE 83 ). Mean dentition formula 31:1:31 (SD = 8.1), mean transverse rows 138 (SD = 20.1, n = 3, AM C.201775, C.201765, C.316303); single central rachidian tooth flanked squarely by 31 half row laterals, 0–4 are inner (SD = 4.98), 12–14 mid (SD = 4.98) and 15–19 outer (SD = 3.58) laterals (n = 5); central tooth with narrow unicuspid mesocone less than half length of basal plate, lower than laterals; inner laterals absent or few ( Figs 83E–F View FIGURE 83 ), mid lateral mesocones generally unicuspid, however, maybe irregularly bicuspidate ( Figs 83E–F View FIGURE 83 ), mid laterals with pointed ectocone; ectocone strong, protrudes at acute angle halfway along the tooth’s length; outer laterals typically with a ‘chisel’ shaped mesocone flanked by small, pointed single ecto and endocones, endocone positioned higher on side of tooth than ectocone, angle of separation of each cone from the mesocone varies ( Fig. 83G View FIGURE 83 ). Jaw located inside front of buccal cavity, orange-brown, arch shaped with ‘shingle’- arranged cone-like rods, ~ 140 rods wide (~ 1.9 mm) by ~ 16 rods deep (mean length = 50 μm, SD = 6, mean width = 12 μm, SD = 3, n = 15); tip bluntly rounded [SK030].

Comparative remarks. Siphonaria denticulata ( atra group, unit 33) is the sister taxon of a clade containing two species, S. opposita (unit 34) and S. plana (unit 35) ( Figs 1–4 View FIGURE 1 View FIGURE 2 View FIGURE 3 View FIGURE 4 ). It differs from these species by COI distances of ≥ 14.8% (unit 35) and ≥ 15.8% (unit 34) (Table S3).

We found eight congeners with sympatric occurrences in SE Australia. For comparisons with S. atra and S. diemenensis refer to comparative remarks under these species. Siphonaria stowae has a smaller, more elongate, paler shell with a strongly offset apex, less prominent ribbing and edge scalloping, a smaller AO, shorter ED larger BC, and shorter SPM. Siphonaria emergens has a smaller, elongate, paler brown shell with a strongly offset apex, less prominent ribbing and edge scalloping. Siphonaria funiculata has a paler, taller shell with less raised ribs broadening to shell edge, a fainter scalloped edge, a larger AO, shorter BD and ED, absent to smaller F1, and larger, drop-like SPM. Siphonaria pravitas sp. nov. has a lower shell with greater raised ribbing and edge scalloping, no BD distal loop, a larger BC, wider F1, and shorter SPM. Siphonaria scabra has a paler shell with finer ribbing and coarser exterior, a longer AO, BC and F1, and a bursal loop. Siphonaria zelandica has a paler, lower shell with less prominent and finer narrower ribbing, a shorter ED and BD, and a short. drop-like SPM.

Menke (1851: 38) referred to the name S. denticulata for a specimen from Mexico. This reference was subsequently considered as a misidentification of S. lecanium var. palmata Carpenter, 1857 ( Carpenter 1857: 183). Hubendick (1946: 40) treated this taxon as a junior synonym and subspecies of S. maura Sowerby I, 1835 .

Erroneous records of ‘ S. denticulata ’ from NZ ( Hutton, 1873: 55) and Mozambique ( Braga, 1956: 7, pl. 1, fig. 2) are herein attributed to S. propria Jenkins, 1983 and possibly S. ferruginea Reeve, 1856 , respectively. We have not located any samples of this species from NZ nor Mozambique in any of the examined museum collections. The treatment of S. denticulata as a synonym or variety of S. diemenensis by several authors is refuted herein and both species are accepted as distinct species. Hedley (1915; 1918; M96) and Angas (1867: 232) incorrectly recorded S. denticulata from Port Jackson (= Sydney) and NSW.

A specimen figured as ‘ S. marza’ in Hubendick (1946: 61, pl. 5, figs 27–28) from Port Jackson [Sydney Harbour]) is herein identified as S. denticulata . McAlpine (1952: 40, 42, fig. 1) misidentified specimens of S. scabra as ‘E llsiphen ’ (sic Ellsiphon ) denticulatus . The RS figured herein ( Figs 20D, G View FIGURE 20 ) matches closely the figure in McAlpine (1952: 43, fig. 1; as Ellsiphon denticulatus ) and that shown in ( Fig. 20F View FIGURE 20 , from Currumbin) closely matches the distal RS figure of ‘ S. currumbinensis ’ from in Hubendick (1955: fig. 4). A specimen from NZ labelled as ‘ S. denticulata ’ in Davey (1998: 117) is herein identified as S. funiculata . Specimens stated to resemble either S. diemenensis , S. denticulata or S. australis by Dayrat et al. (2014) as ‘unit 7’ are herein identified as S. diemenensis . By contrast, the figured specimens of ‘ atra group, unit 33’ in Dayrat et al. (2014: fig. 5J, K) are morphologically consistent with S. denticulata .

Distribution and habitat. Endemic to temperate eastern and southeastern Australia, from Keppel Bay, Qld, south to Cape Conran, Vic, and Bridport, Tas ( Fig. 16 View FIGURE 16 ). In this study found in sheltered to moderately exposed sites on hard substrates (e.g., rocky shores, wooden pilings); upper and mid littoral levels; home scars prominent.